Disentangling the components of pollen limitation in a widespread herb with gametophytic self-incompatibility.

PREMISE: Seed production is frequently limited by the receipt of insufficient or low-quality pollen, collectively termed "pollen limitation" (PL). In taxa with gametophytic self-incompatibility (GSI), incompatible pollen can germinate on stigmas but pollen tubes are arrested in styles. This allows for estimates of pollen performance before, during, and after self-recognition, as well as insight into the factors underlying pollen quality limitation in GSI taxa. METHODS: We scored pollen performance following self and outcross pollinations in Argentina anserina to identify the location of self-recognition and establish the relationship between pollen tubes and seed production. We then estimated quantity and quality components of PL from >3300 field-collected styles. We combined our results with other studies to test the prediction that low pollen quality, but not quantity, drives higher PL in self-incompatible (SI) taxa than in self-compatible taxa (SC). RESULTS: Self and outcross pollen germinated readily on stigmas, but 96% of germinated self-pollen was arrested during early tube elongation. Reproduction in the field was more limited by pollen quality than by quantity, and pollen failure near the location of self-recognition was a stronger barrier to fertilization than pollen germination. Across 26 taxa, SI species experienced stronger pollen quality, but not quantity, limitation than SC species. CONCLUSIONS: Evaluating pollen performance at multiple points within pistils can elucidate potential causes of pollen quality limitation. The receipt of incompatible pollen inhibits fertilization success more than insufficient pollen receipt or poor pollen germination in A. anserina. Likewise, pollen quality limitation drives high overall PL in other SI taxa.

A test of Sensory Drive in plant-pollinator interactions: heterogeneity in the signalling environment shapes pollinator preference for a floral visual signal.

Sensory Drive predicts that habitat-dependent signal transmission and perception explain the diversification of communication signals. Whether Sensory Drive shapes floral evolution remains untested in nature. Pollinators of Argentina anserina prefer small ultraviolet (UV)-absorbing floral guides at low elevation but larger guides at high. However, mechanisms underlying differential preference are unclear. High elevation populations experience elevated UV irradiance and frequently flower against bare substrates rather than foliage, potentially impacting signal transmission and perception. At high and low elevation extremes, we experimentally tested the effects of UV light (ambient vs reduced) and floral backgrounds (foliage vs bare) on pollinator choice for UV guide size. We examined how different signalling environments shaped pollinator-perceived flower colour using visual system models. At high elevation, pollinators preferred locally common large UV guides under ambient UV, but lacked preference under reduced UV. Flies preferred large guides only against bare substrate, the common high elevation background. Ambient UV amplified contrast of large UV guides with floral backgrounds, and flowers contrasted more with bare ground than foliage. Results support that local signalling conditions contribute to pollinator preference for a floral visual signal, a key tenet of Sensory Drive. Components of Sensory Drive could shape floral signal evolution in other plants spanning heterogeneous signalling environments.

Elevational divergence in pigmentation plasticity is associated with selection and pigment biochemistry.

Phenotypic plasticity is predicted to evolve in environmentally variable habitats, or those experiencing a high frequency of strong selection. The evolution of plasticity may however be constrained by costs or physiological limitations. In flowers, UV-absorbing pigmentation ameliorates UV damage to pollen, and is linked with elevated UV exposure. Whether plasticity contributes to this pattern remains unclear. Petals of Argentina anserina have larger UV-absorbing petal areas at high elevations where they experience higher and more variable UV exposure than low elevations. We measured UV-induced pigmentation plasticity in high- and low-elevation populations (hereafter, "high," "low"), and selection on pigmentation via male fitness. We dissected UV pigment biochemistry using metabolomics to explore biochemical mechanisms underlying plasticity. High displayed positive UV-induced pigmentation plasticity but low lacked plasticity. Selection favored elevated pigmentation under UV in high, supporting adaptive plasticity. In high, UV absorption was conferred by flavonoids produced in one flavonoid pathway branch. However, in low, UV absorption was associated with many compounds spanning multiple branches. Elevated plasticity was thus associated with reduced pigment diversity. These results are consistent with adaptive floral pigmentation plasticity in more extreme and variable environments. We discuss how biochemical underpinnings of pigmentation may permit or constrain the evolution of pigmentation plasticity.